lemon on Troyer Citrange Root bud-union and rootstock disorder of Troyer citrange with Eureka tops under study in effort to identify cause

I.. G. Weathers, E. C. Calavan, J. M. Wallace, and D. W. Christiansen

A serious disorder in trees of Eureka dry, and stained light brown, but lacked tions. Although there was no conclusive lemon on Troyer citrange-a hybrid of the usual odor of dry root rot. evidence of a causal relationship between the trifoliate and the navel or- In some trees the necrosis involved the the bud-union disorder and the necrotic ange-rootstock was discovered in 1954 entire rootstock, while in others it OC- lesions: it was observed that the bud- when all the Eureka tops in one of the curred as localized lesions or in longitu- union disorder of Eureka lemon on experimental plots in Ventura County dinal strips. The lesions did not extend Troyer citrange frequently occurs in the turned yellow, set a heavy crop of fruit, into the Eureka lemon portion of the absence of rootstock lesions. With this and declined markedly. trunk. Both old-line and nucellar-line combination no lesions were found with- Examination revealed a disturbance at Eureka on Troyer in this - out a conspicuous bud-union disorder. the bud union, with retarded growth, de- ing showed the disease, but the same tops Cook nucellar-line Eureka lemons top- terioration, and gumming of the Troyer on other rootstocks were not similarly worked on Valencia sweet orange on citrange. Bark and wood necrosis, ac- affected. Troyer citrange rootstocks in one grove companied by some gumming and resem- A survey of other plantings of lemon in Ventura County were found declining bling dry root rot, had developed in the on Troyer citrange in Ventura and Santa severely three years after topworking. rootstocks. The affected wood was firm, Barbara counties revealed that many of Concluded on next page the 3- to 4-year-old trees of Frost, UCLA, Cook, and Hughes nucellar-line, and of Allen, Hughes, and Cascade old-line Eureka lemons, propagated on Troyer citrange, were declining or had died. Other cases have been observed recently in Orange County. The bud unions of affected trees were noticeably abnormal, with the lemon scion having overgrown the stock and the bark of the Troyer often having dried and adhered to the wood. A narrow dark- brown line in the bark and on the cam- bial face of the wood at the bud union suggested some form of incompatibility. Treatment with iodine showed starch depletion in the Troyer rootstocks, which is indicative of girdling near the bud union of a sufficient degree to restrict severely the translocation of carbohy- drates to the roots. Wood necrosis had developed in the Troyer rootstocks of only a few trees and appeared to be a secondary symptom associated with fun- gus invasion of tissues weakened by girdling. Some trees without extensive wood necrosis have made a partial re- covery. There was considerable variation in symptom expression on trees growing under different environmental condi-

A. Disorder of Troyer citrange rootstock with Eureka lemon top, showing gumming at the bud union and retarded growth of the rootstock. B. Cross section of lroyer citrange rootstock, showing the developing necrosis. C. localized lesion on Troyer citrqnge rootstock. D. Longitudinal section through bud union of diseased Troyer citrange rootstock with Eureka lemon scion. Note the sharp line of demarcation at the union and the discolored wood of the rootstock. (Photographs A, B, and D by R. G. Platt; C by E. L. Yiddleham.)

CALIFORNIA AGRICULTURE, NOVEMBER, 195 5 11 LEMON and possibly all old-line Eureka lemons carry a transmissible factor, possibly a Continued from preceding page virus, which retards the growth of nucel- Lesions had developed in the Troyer lar Eureka lemons. The retardation of rootstocks of some trees, but no brown growth of nucellar-line Eureka lemons line or marked overgrowth was found at after graft-inoculation from old-line Eu- either the lemon-orange or the orange- rekas does not appear to be related to citrange unions, as was the case where the disorder of Eureka lemon on Troyer Eureka lemon was propagated directly citrange. The virus or causal factor of on Troyer citrange. However, the bark the former is not present in nucellar-line of the Troyer rootstock was very thin Eureka lemons; whereas the factor re- and dry and adhered very tightly to the sponsible for the Eureka-Troyer disor- wood. Tests with iodine indicated low der is present in all the nucellar-line starch content in the roots. Eurekas tested. Fungi of several genera were isolated (Photograph by K. L. Middleham) Trees of Frost and UCLA nucellar-line from the bark and wood of diseased Bud-union disorder of nucellar Eureka lemon Eureka on some rootstocks other than Troyer citrange rootstocks from several on rootstock one year after Troyer citrange or trifoliate orange are budding. Bark removed to show groove and locations in Ventura County. Fungi com- dark line at the union. Rule is in inches. retarded in growth only after being monly found were Alternariu spp., Fu- graft-inoculated from old-line Eureka sarium spp., Penicillium spp., and a sources, but the same nucellar lines when fungus tentatively identified as Gliocla- lar-line Eureka lemon on rootstocks of budded on Troyer citrange subsequently dium roseum Bainier. These isolates have trifoliate orange, Po'mirus trifoliata develop the Troyer rootstock disorder not yet been fully tested by inoculation (Linn.) Raf., sometimes within a year without inoculation. If a virus is involved into Troyer citrange, but from previous after budding, have recently been ob- in the latter disorder, it must be present tests it is known that they are ordinarily served. The disorder of Eureka lemon generally in nucellar Eurekas or, as is present on as saprophytes or weak on trifoliate orange reveals a bud-union somewhat less likely, seed-transmitted parasites of weakened tissues. Attempts condition very similar to that observed and carried latently in Troyer citrange to isolate spp. from the on trees of Eureka lemon on Troyer cit- seedlings. It is also known that trifolate diseased tissues and from the soil ad- range rootstocks. A young Messina-type orange rootstocks having certain old-line jacent to the Troyer rootstocks were un- lemon on trifoliate orange in Santa Bar- Eureka lemon tops show symptoms of successful. bara County has also developed a bud- exocortis. However, exocortis is not The cause of this disorder of Troyer union disturbance and exocortis. Wood necessarily the cause of either the bud- citrange rootstock has not been deter- necrosis has not been observed in any union trouble or the necrosis of Troyer mined. However, the following points of the diseased trifoliate orange root- citrange. were noted: 1, Eureka is the only lemon stocks. Experiments aimed at identifying the variety known to be associated with this agent or agents responsible for the Troy- disorder. Lisbon lemon strains and or- Possible Causal Agents er citrange rootstock disorder are now anges on Troyer citrange rootstocks have in progress. The Troyer citrange rootstock dis- not thus far been similarly affected. 2, L. G. Weathers is Assistant Plant Pathologist, Until symptoms appear two or more order may be caused by a virus or by a University of California, Riverside. years after planting, the trees are very form of incompatibility between Eureka E. C. Calavan is Associate Plant Pathologist, vigorous and apparently healthy. The lemon and Troyer citrange. The occur- University of California, Riverside. most vigorous trees are often the first to rence of rootstock lesions in trees having .I.M. Wallace is Plant Pathologist, Uniuer- deteriorate. 3, Soil type, soil fumigation Valencia sweet orange interstocks dis- sity of California, Riverside. with DD, and irrigation practices ap- favors direct incompatibility as the D. W. Christiansen is Senior Laboratory Technician, University of California, Riverside. pear to have little or no influence on the cause of this symptom, since the Valen- cia interstock separating the Eureka Dr. L. D. Batchelor, Dr. W. P. Bitters, G. E. appearance of this disorder. 4, Many Goodall, R. G. Platt, and the late F. A. White, trees become diseased before developing lemon from the Troyer citrange is com- University of California, co-operated in the wood rot or any other evidence of fungus patible with both. This indicates that the studies reported here, as did Citrus Field Re- activity. Since the fungi thus far isolated Eureka lemon may carry a virus that search, Inc., Limoneira Company, Rancho seriously affects Troyer citrange. Fur- Sespe, Utt Development Company, Ltd., Ven- from diseased roots appear to be second- tura Coastal Lemon Company, and individual ary invaders, it is very doubtful that thermore, inasmuch as nucellar lines of growers. Eureka on Troyer citrange react in the fungi are the primary .cause. 5, Several The above progress report is based on Re- cases of failure of old-line and nucel- same way as trees of old-line Eureka on search Project Nos. 1383 and 1544. this rootstock, any virus present must be either seed-transmissible or very readily transmissible by mechanical means or by a vector. Other studies have shown that some

A. Four-year-old nucellar Eureka lemon on Troyer citrange. A flop of bark has been peeled downward across the union to show the dark line on the cambial faces of the wood and bark. B. Four-year-old nucellar Eureka lemon on Troyer citrange. The ,bark across the union was cut away. A groove was chiseled across the union and the wood treated with iodine. Note color reaction for starch in the Eureka lemon wood above the union (arrow) and lack of color reac- tion in the Troyer citrange wood. (Photograph A by K. L. Middleham; B by R. Q. Platt.)

12 CALIFORNIA AGRICULTURE, NOVEMBER, 1955